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Reappraisal of Austrosaurus Mckillopi Longman, 1933 from the Allaru

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Reappraisal of Austrosaurus Mckillopi Longman, 1933 from the Allaru 1 1 Reappraisal of Austrosaurus mckillopi Longman, 1933 from the 2 Allaru Mudstone of Queensland, Australia’s first named 3 Cretaceous sauropod dinosaur 4 5 STEPHEN F. POROPAT, JAY P. NAIR, CAITLIN E. SYME, PHILIP D. MANNION, 6 PAUL UPCHURCH, SCOTT A. HOCKNULL, ALEX G. COOK, TRAVIS R. TISCHLER 7 and TIMOTHY HOLLAND 8 9 POROPAT, S. F., NAIR, J. P., SYME, C. E., MANNION, P. D., UPCHURCH, P., HOCKNULL, S. A., 10 COOK, A. G., TISCHLER, T.R. & HOLLAND, T. 201#. Reappraisal of Austrosaurus mckillopi 11 Longman, 1933 from the Allaru Mudstone of Queensland, Australia’s first named Cretaceous 12 sauropod dinosaur. Alcheringa XX, XX–XX. ISSN ###. 13 14 Austrosaurus mckillopi Longman, 1933 was the first Cretaceous sauropod reported from 15 Australia, and the first Cretaceous dinosaur reported from Queensland (northeast Australia). 16 This sauropod taxon was established on the basis of several fragmentary presacral vertebrae 17 (QM F2316) derived from the uppermost Lower Cretaceous (upper Albian) Allaru Mudstone, 18 at a locality situated 77 km west-northwest of Richmond, Queensland. Prior to its rediscovery 19 in 2014, the type site was considered lost after failed attempts to relocate it in the 1970s. 20 Excavations at the site in 2014 and 2015 led to the recovery of several partial dorsal ribs and 21 fragments of presacral vertebrae, all of which clearly pertained to a single sauropod dinosaur. 2 22 The discovery of new material of the type individual of Austrosaurus mckillopi, in tandem 23 with a reassessment of the material collected in the 1930s, has facilitated the rearticulation of 24 the specimen. The resultant vertebral series comprises six presacral vertebrae—the 25 posteriormost cervical and five anteriormost dorsals—in association with five left dorsal ribs 26 and one right one. The fragmentary nature of the type specimen has historically hindered 27 assessments of the phylogenetic affinities of Austrosaurus, as has the fact that these 28 evaluations were often based on a subset of the type material. The reappraisal of the type 29 series of Austrosaurus presented herein, on the basis of both external morphology and 30 internal morphology visualised through CT data, validates it as a diagnostic titanosauriform 31 taxon, tentatively placed in Somphospondyli, and characterised by the possession of an 32 accessory lateral pneumatic foramen on dorsal vertebra I (a feature which appears to be 33 autapomorphic) and by the presence of a robust ventral midline ridge on the centra of dorsal 34 vertebrae I and II. The interpretation of the anteriormost preserved vertebra in Austrosaurus 35 as a posterior cervical has also prompted the re-evaluation of an isolated, partial, posterior 36 cervical vertebra (QM F6142, the “Hughenden sauropod”) from the upper Albian Toolebuc 37 Formation (which underlies the Allaru Mudstone). Although this vertebra preserves an 38 apparent unique character of its own (a spinopostzygapophyseal lamina fossa), it is not able 39 to be referred unequivocally to Austrosaurus and is retained as Titanosauriformes indet. 40 Austrosaurus mckillopi is one of the oldest known sauropods from the Australian Cretaceous 41 based on skeletal remains, and potentially provides phylogenetic and/or palaeobiogeographic 42 context for later taxa such as Wintonotitan wattsi, Diamantinasaurus matildae and 43 Savannasaurus elliottorum. 44 3 45 Stephen F. Poropat [[email protected]; [email protected]] (corresponding 46 author), Department of Chemistry and Biotechnology, Swinburne University of Technology, 47 John St, Hawthorn, Victoria 3122, Australia and Australian Age of Dinosaurs Museum of 48 Natural History, Lot 1 Dinosaur Drive, PO Box 408, Winton, Queensland 4735, Australia; 49 Jay P. Nair [[email protected]; [email protected]] School of Biological Sciences, The 50 University of Queensland, St Lucia, Queensland 4072, Australia; Caitlin E. Syme 51 [[email protected]] School of Biological Sciences, The University of 52 Queensland, St Lucia, Queensland 4072, Australia; Philip D. Mannion 53 [[email protected]] Department of Earth Science and Engineering, Imperial 54 College London, South Kensington Campus, London SW7 2AZ, United Kingdom; Paul 55 Upchurch [[email protected]] Department of Earth Sciences, University College 56 London, Gower Street, London WC1E 6BT, United Kingdom; Scott A. Hocknull 57 [[email protected]] Geosciences, Queensland Museum, 122 Gerler Rd, Hendra, 58 Queensland 4011, Australia; Alex G. Cook [[email protected]] School of Earth 59 Sciences, The University of Queensland, St Lucia, Queensland 4072, Australia; Travis R. 60 Tischler [[email protected]] Australian Age of Dinosaurs Museum of Natural 61 History, Lot 1 Dinosaur Drive, PO Box 408, Winton, Queensland 4735, Australia; Timothy 62 Holland [[email protected]] Kronosaurus Korner, 91 Goldring St, Richmond, 63 Queensland 4822, Australia. Received XX.X.201X; revised XX.X.201X; accepted XX.X.201X. 64 65 Key words: Austrosaurus; Dinosauria; Sauropoda; Titanosauriformes; Australia; Cretaceous; 66 Gondwana. 67 4 68 SAUROPOD dinosaur fossils were not reported from Australia until 1926, despite the 69 fact that the first discovery of an Australian sauropod specimen was made in 1913. This 70 incomplete humerus (QM F311)—found in Cretaceous strata near the town of Blackall in 71 Queensland—was not determined to pertain to a dinosaur, let alone a sauropod, until 1980 72 (Molnar 2001b). The first Australian sauropod specimen that was recognised as such was 73 discovered in 1924 near the town of Roma, Queensland. This partial skeleton (QM F1659) 74 was designated the type of Rhoetosaurus brownei and remains one of only two Jurassic 75 sauropod specimens known from Australia (Longman 1926, 1927a, b, 1929, Thulborn 1985, 76 Rich & Vickers-Rich 2003, Nair & Salisbury 2012). The other, a distal caudal vertebra 77 (UWA 82468), was found in the mid-1970s in Middle Jurassic rocks near Geraldton, Western 78 Australia (Long 1992). 79 Cretaceous sauropod remains are only known from three Australian states. Footprints 80 found in the Broome Sandstone (Valanginian–Barremian) of the Dampier Peninsula 81 constitute the only evidence of Cretaceous sauropods in Western Australia (Thulborn et al. 82 1994, Thulborn 2012, Salisbury et al. 2017), whereas opalised sauropod teeth (AM F66769, 83 AM F66770) from the Griman Creek Formation (middle Albian) of Lightning Ridge are all 84 that has been reported from New South Wales (Molnar & Salisbury 2005, Molnar 2011b). In 85 Queensland, the Griman Creek Formation has yielded two possible sauropod elements: a 86 fragmentary right ischium (QM F54817) and a possible vertebral fragment (QM F11043; 87 Molnar 2011b). 88 The Eromanga Basin in Queensland has produced the overwhelming majority of 89 Australia’s sauropod skeletal remains. Rare specimens have been described from the 90 Toolebuc Formation and the Allaru Mudstone, both of which constitute upper Albian marine 5 91 deposits (Molnar & Salisbury 2005). However, the bulk of the known sauropod remains from 92 the Eromanga Basin derive from the youngest Cretaceous unit preserved therein, the 93 uppermost Albian–lower Turonian Winton Formation (Coombs & Molnar 1981, Molnar & 94 Salisbury 2005, Greentree 2011, Bryan et al. 2012, Tucker et al. 2013, Tucker 2014), which 95 evinces the existence of a vast floodplain. To date, three sauropod taxa have been reported 96 from the Winton Formation: the non-titanosaurian somphospondylan Wintonotitan wattsi 97 (Hocknull et al. 2009, Poropat et al. 2015a); and the titanosaurs Diamantinasaurus matildae 98 (Hocknull et al. 2009, Poropat et al. 2015b, 2016) and Savannasaurus elliottorum (Poropat et 99 al. 2016). However, the first Cretaceous sauropod, and indeed the first Cretaceous dinosaur, 100 reported from Queensland was not from the Winton Formation: it was found in the Allaru 101 Mudstone, and bears the name Austrosaurus mckillopi (Longman 1933). 102 ===PLEASE INSERT FIGURE 1=== 103 The type and only known specimen of Austrosaurus mckillopi was found in August 104 1932 on Clutha sheep station, 77 km west-northwest of Richmond, Queensland (Fig. 1). 105 Clutha overseer Henry Burgoyne Wade (Fig. 2A) found fragments of fossilised bone near the 106 southwest corner of Whitewood Paddock. He showed these to the station manager, Harley 107 John McKillop (Fig. 2B), who contacted his brother, Dr Martin Joseph McKillop (Fig. 2C). 108 Shortly afterwards, M. J. McKillop travelled from Brisbane to Clutha, helped H. B. Wade and 109 H. J. McKillop to recover additional specimens, and sent a sketch of one to Heber Albert 110 Longman (Fig. 2D), then director of the Queensland Museum. Longman requested that the 111 specimens be sent to Brisbane, and they arrived in January 1933. By March 1933, Longman 112 had correctly determined that the bones derived from a sauropod dinosaur and made them the 113 type specimen of Austrosaurus mckillopi, honouring M. J. McKillop with the species epithet 6 114 (Longman 1933). A sign was erected by H. B. Wade and H. J. McKillop at the type site to 115 demarcate its significance (Fig. 2E). 116 ===PLEASE INSERT FIGURE 2=== 117 In his original description of Austrosaurus mckillopi, Longman only mentioned three 118 specimens (and figured two), all of which were presacral vertebrae (catalogued as QM 119 F2316). It is probable that Longman received more than three specimens in the initial 120 shipment from Clutha, given that contemporary newspaper articles stated that six vertebrae 121 were preserved (Anonymous 1933b, a). However, only on the three specimens described by 122 Longman is there evidence of “repeated soakings in shellac solution” (Longman 1933, p. 123 132). Correspondence between H. J. McKillop and Longman confirms that a second 124 shipment of specimens was dispatched from Clutha to the Queensland Museum in June 1933. 125 Molnar (2001b, p. 141, 2010, p. 423, 2011a, p. 332) and Molnar & Salisbury (2005, p. 456) 126 suggested that this second shipment comprised five large and more than ten small pieces. 127 However, in his letter to Longman on 23/05/1933, H. J.
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